ASSESSING POTENTIAL ECOLOGICAL EFFECTS OF MUTE SWAN (CYGNUS OLOR) EXPANSION IN NORTHEASTERN NORTH AMERICA Kristen Bell Travis MS and Erik Kiviat PhD Hudsonia P.O. Box 5000 Annandale NY 12504 Prepared for Grant & Lyons LLP Rhinebeck NY 12 October 2017 INTRODUCTION Mute swan (Cygnus olor) management and proposed future management in North America have been controversial. Conservation controversies can benefit from an impartial and transparent evaluation of available scientific evidence, including its gaps and uncertainties (Redpath et al. 2013). Conservation practice is often guided by myth rather than evidence (Sutherland et al. 2004), or by evidence drawn from flawed studies (e.g., Allen et al. 2013). This paper reviews what we know of mute swan ecology relative to its potential effects in northeastern North America, to assess the scientific underpinnings of proposed management goals and methods. In this review we draw extensively on ecological studies of mute swan in both its native and introduced ranges, in order to evaluate claims of this large aquatic herbivore’s impacts on aquatic vegetation (including food resources for other waterfowl), and its competitive or agonistic (aggressive) behavior toward other species (and potential displacement of nesting waterbirds). These claims have formed the basis of management plans in many U.S. states. We summarize relevant aspects of mute swan biology and ecology, including habitat use, seasonal flocking, territoriality, and diet. We also examine the larger context of aquatic vegetation dynamics, including the roles of a demanding and changeable physical environment, herbivory, and anthropogenic stressors in the maintenance or decline of these communities. To assess the potential effects of mute swans on natural communities and organisms of conservation concern, we rely as much as possible on robust experimental and observational studies. Where case studies or single observations are discussed, we identify them as such. We discuss potential interactions between mute swans and other expanding native and nonnative species, and touch on ecosystem processes that could be affected by swans. We outline possible effects of swans on humans, identify important unanswered questions, discuss factors that limit mute swan populations, and review the effectiveness of different management actions and potential barriers to successful management. Hudsonia is not supporting or opposing mute swan management in general, or any particular management proposal or plan. We are examining the scientific evidence and analysis that are cited to support swan management in many states. HISTORY Mute swans are considered native to parts of north and central Eurasia, although a long history of semi-domestication has made distinctions between their natural and introduced range difficult (Scott 1972). Swans have played an important role in mythology and art around the world, and the “brilliant creatures” of W.B. Yeats continue to be valued for their beauty. They have also been harvested extensively for meat and skins. Historically, swans of all species (including mute swans) were hunted at any time of year, and sometimes rounded up and killed by the hundreds when flightless during molt. However, the mute swan is unique in its semi-domestication (i.e., 2 with a history of being bred and raised in captivity but genetically indistinguishable from wild birds) from very early times. In England, mute swan ownership was a mark of social standing denoted by marking adult birds; any unmarked swans were (and still are) property of the Crown. Mute swan was prized for its meat, often serving as the centerpiece of a banquet, and also as an ornament on estates (Scott 1972). Populations in many European countries were very small or extirpated by the early 1900s. European populations have greatly increased in abundance since the 1930s (Scott 1972) and have both expanded in range and increased in abundance since the 1970s (Weiloch 1991), more than doubling between the late 1980s and early 2000s (Gayet et al. 2014). This growth may be due to some combination of the swan’s protected status (in many countries dating from the early–mid-20th century), wetland eutrophication (increased fertility), warmer winters, the creation of new habitat, and increased winter food availability due to the intensification of agriculture and human handouts (Gayet et al. 2014). Mute swans have shown great plasticity in migration distance, location of breeding, molting, and wintering areas, territoriality, and use of food resources, which has apparently enabled them to expand into areas with warmer winters (where they tend to become less migratory) and into agricultural and urban landscapes (where they depend to a variable degree on crops and human handouts for winter food). The eastern U.S. population probably originated from multiple introductions to four areas: Dutchess County, New York; the southern shore of Long Island, Suffolk County, New York; the eastern shore of New Jersey (Monmouth and Ocean counties); and Boston—during the first three decades of the twentieth century (Ciaranca et al. 1997; Teale 2011). The swans were originally imported from Europe to decorate estates; their wings were generally clipped to prohibit escape. They were first documented breeding in the wild in the 1920s in New York (Teale 2011) and in the 1940s-1960s in other eastern states (Ciaranca et al. 1997). The U.S. eastern seaboard population grew from approximately 6,000 in 1986 to 14,000 in 2002, then declined to 9,000 in 2011 due to management. Naturalized and expanding populations have also been established along the Great Lakes, in the Pacific Northwest, and in Australia, New Zealand, South Africa, and Japan (Ciaranca et al. 1997). POPULATION IN THE NORTHEAST Regional growth rates in the Atlantic Flyway between 1986 and 1999 ranged from 43% in New England (mostly in Massachusetts, Rhode Island, and Connecticut) to 62% in the upper Mid- Atlantic states (mostly in New York, New Jersey, and Pennsylvania) to 1271% in the Chesapeake Bay Region (Maryland and Virginia; Costanzo et al. 2015). At more local scales, mute swan populations may double every five years in the initial growth phase after establishment, but then population growth slows after 25-30 years due to density dependence (Ellis & Elphick 2007). Mute swans have been actively but not consistently managed (through egg addling and/ or culling or relocating adult birds) in several Atlantic Flyway states: Maryland, Delaware, Rhode Island, New York, and Vermont (State of Rhode Island 2006). Surveys in 2008 and 2011 recorded approximately 2,600-2,900 swans in New England (mostly in Rhode Island, Connecticut, and Massachusetts), 2,800-3,900 in New York and New Jersey, and approximately 3,000 in Ontario, mostly along the Lower Great Lakes (Meyer et al. 2012; Costanzo et al. 2015). Chesapeake Bay had over 4,400 swans in 1999 but management reduced survey results to approximately 300 by 2011 (Costanzo et al. 2015). 3 In the Northeast, mute swans occur at the highest densities along coastal and Great Lakes shorelines and estuaries, but are increasingly common inland in freshwater ponds, rivers, and wetlands (State of Rhode Island 2006; Swift et al. 2013; Costanzo et al. 2015), a trend also noted in Ontario (Meyer et al. 2012) and Sweden (Berglund et al. 1963). At some coastal locations in Rhode Island, Connecticut, New York, and adjacent areas, nesting mute swans may have reached the local carrying capacity (Ellis & Elphick 2007; Swift et al. 2013). Breeding Bird Atlas data show a substantial expansion in distribution inland in New York between 1985 and 2005 (Swift et al. 2013). SEASONAL HABITAT USE, FLOCKING, AND TERRITORIALITY Habitat requirements of mute swans are similar year-round, and ideally consist of open, shallow water (20-45 cm deep) with abundant aquatic macrophytes, especially submergent plants and macroalgae (also referred to as submerged aquatic vegetation [SAV]). They may be found in a variety of saline to freshwater, natural or constructed, estuaries, wetlands, rivers, lakes, and ponds that have these characteristics (Ciaranca et al. 1997), and they are more likely to be found near urban land cover (Weaver et al. 2012). Mute swans are either year-round residents or short- distance migrants, and their behavior and habitat use depend on swan age and breeding status as well as season and habitat condition. Understanding the (often complex) linked consequences of these factors is prerequisite for understanding the swans’ potential effects on habitats and other species. Breeding territories and both summer and winter flocking sites may be in the same or adjacent areas, or be separated by dozens to hundreds of kilometers. Breeding pairs establish territories in late winter or early spring. Territory size and spacing vary widely depending on habitat quality and swan density. The area of defended territory generally corresponds to the area the pair uses, and can vary from, e.g., 0.2 to 4.8 ha (based on two North American and several European studies; Ciaranca et al. 1997 [one hectare = 2.47 acres]). They also occasionally nest in colonies (nests spaced 10-30 m apart) on coastal islands (Scott 1972; Ciaranca et al. 1997). Territory size calculated as the spacing between nests varies even more: mean areas of 4.5, 500, and 2,000 ha in three sites with suitable habitat in their native range (Scott 1972); minimum areas of 0.8 and 1.9 ha in Rhode Island; and a mean area of 3.9 ha in Michigan (Ciaranca et al. 1997). Pairs often defend their territories, primarily by using threat displays to chase away other birds, but occasionally attacking with wings or bill. Males are generally more aggressive than females, and levels of aggression vary greatly by individual or pair. The pair will usually remain on the territory until food is depleted or water freezes. Meanwhile, nonbreeding swans—including juveniles, first-year birds, and a large proportion of adults (e.g., 71% of adults were nonbreeders in one U.K.